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370 Biotechnological Approaches for Pest Management and Ecological Sustainability<br />

In this process, no crystal solubilization and protoxin-toxin conversion is necessary in the<br />

insect midgut. Increased toxicity has been observed in some herbivorous insects when fed<br />

with the toxin instead of the pro-toxin, suggesting that such insects lack the appropriate<br />

enzyme composition or pH necessary to convert pro-toxin to toxin (Moar et al., 1995). The<br />

truncated and activated toxins expressed in transgenic plants in high doses kill the insects<br />

much faster than Bt toxins applied as sprays, which need to be activated by enzymes in the<br />

insect midgut. While Bt sprays might affect all the insects in an ecosystem wherever they<br />

are sprayed, the toxins from the transgenic plants affect only the insects that ingest the toxin<br />

from the plants through feeding or indirectly through the insect host or prey. Therefore,<br />

toxicity results and development of resistance to Bt formulations cannot be substituted for<br />

the toxins expressed in transgenic plants. Development of resistance and insect sensitivity<br />

to Bt toxins and other novel genes is not properly understood, and such information is<br />

important in the context of deployment of transgenic plants for pest management (Sharma<br />

and Ortiz, 2000). Since plants expressing only the Bt cry proteins have been released for<br />

cultivation, this chapter will largely focus on issues related to development of resistance to<br />

Bt Cry proteins and strategies for resistance management for sustainable crop production.<br />

Tremendous progress had been made in developing insect-resistant transgenic plants<br />

with genes from Bt. Genetically modifi ed plants with resistance to insects were developed<br />

in the mid-1980s (Barton, Whiteley, and Yang, 1987; Vaeck et al., 1987), and since then, there<br />

has been rapid progress in developing transgenic plants with insect resistance in several<br />

crops (Hilder and Boulter, 1999; Sharma, Sharma, and Crouch, 2004). Transgenic crops<br />

are now being grown on over 100 million ha worldwide (James, 2007). The success of Bt<br />

transgenic crops has far exceeded expectations, and does not preclude development of<br />

resistance to Bt toxins expressed in transgenic plants in the future. Key factors delaying<br />

development of resistance in insect populations to Bt crops include refuges of non-Bt host<br />

plants that enable survival of susceptible individuals, low resistance allele frequencies,<br />

recessive inheritance of resistance to Bt toxins, fi tness costs associated with resistance, and<br />

the performance of resistant populations on non-Bt hosts. The relative importance of these<br />

factors may vary among different crops and cropping systems. Widespread and prolonged<br />

cultivation of Bt transgenic crops represents the largest adoption of insect-resistant crops<br />

over time and space (Ferre and Van Rie, 2002). Before large-scale deployment of transgenic<br />

crops for pest management was undertaken, many scientists predicted development of<br />

resistance to transgenic crops at a fast rate, a view re-inforced by selection for resistance to<br />

Bt Cry proteins under laboratory conditions. However, monitoring of insects in regions<br />

with high adoption of Bt crops has not yet led to detection of resistance in fi eld populations<br />

of European corn borer, Ostrinia nubilalis (Hubner), pink bollworm, Pectinophora gossypiella<br />

(Saunders), corn earworm, Helicoverpa zea (Boddie), and tobacco budworm, Heliothis virescens<br />

(Fab.) in the United States; and Helicoverpa armigera (Hubner) in China, India, and<br />

Australia. Despite expectations that insect pests would rapidly evolve resistance to transgenic<br />

crops, increases in the frequency of resistance alleles caused by exposure to Bt crops<br />

in fi eld have not yet been documented (Tabashnik et al., 2003). In laboratory and greenhouse<br />

tests, several strains with resistance to Bt toxins have been selected. However, strains<br />

with 70- to 10,000-fold resistance to Bt toxins in the artifi cial diet failed to complete development<br />

on Bt transgenic crops. Resistance to Bt appears to be costly and there is a rapid<br />

decline of resistance in populations collected from greenhouses and then maintained in<br />

the laboratory without selection.<br />

With the development of resistance to Bt toxins, the value of transgenic crops will diminish<br />

greatly due to lower sensitivity of the target pest to the Bt toxins. One of the consequences<br />

of such a development would be that the farmers would resort to large-scale application of

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